Chlorophyta
Chlorophyta | |
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"Siphoneae" from Ernst Haeckel's Kunstformen der Natur, 1904 | |
Scientific classification | |
Kingdom: | Plantae |
Division: | Chlorophyta Reichenbach, 1828, emend. Pascher, 1914, emend. Lewis & McCourt, 2004[1][2][3] |
Classes[4] | |
Synonyms | |
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Chlorophyta is a division of green algae, informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to a highly paraphyletic group of all the green algae within the green plants (Viridiplantae) and thus includes about 7,000 species[6][7] of mostly aquatic photosynthetic eukaryotic organisms. In newer classifications, it refers to one of the two clades making up the Viridiplantae, which are the chlorophytes and the streptophytes. The clade Streptophyta consists of two divisions, the Charophyta and the Embryophyta.[8][9] In this sense the Chlorophyta includes only about 4,300 species.[4] Like the land plants (bryophytes and tracheophytes), green algae contain chlorophyll a and chlorophyll b and store food as starch[6] in their plastids.
The division contains both unicellular and multicellular species. While most species live in freshwater habitats and a large number in marine habitats, other species are adapted to a wide range of environments. Watermelon snow, or Chlamydomonas nivalis, of the class Chlorophyceae, lives on summer alpine snowfields. Others live attached to rocks or woody parts of trees. Monostroma kuroshiensis, an edible green alga cultivated worldwide and most expensive among green algae, belongs to this group. Some lichens are symbiotic relationships between fungi and green algae.
Members of the Chlorophyta also form symbiotic relationships with protozoa, sponges, and cnidarians. All are flagellated,[10] and these have an advantage of motility. Some conduct sexual reproduction, which is oogamous or isogamous.
Ecology
Species of Chlorophyta (treated as what is now considered one of the two main clades of Viridiplantae) are common inhabitants of marine, freshwater and terrestrial environments.[11][12] Several species have adapted to specialised and extreme environments, such as deserts, arctic environments, hypersaline habitats, marine deep waters and deep-sea hydrothermal vents. [13][14][15] Some groups, such as the Trentepohliales are exclusively found on land.[16] Several species of Chlorophyta live in symbiosis with a diverse range of eukaryotes, including fungi (to form lichens), ciliates, forams, cnidarians and molluscs. [12] Some species of Chlorophyta are heterotrophic, either free-living or parasitic.[17][18] Two common species of the heterotrophic green alga Prototheca are pathogenic and can cause the disease protothecosis in humans and animals.[19]
Classifications
Characteristics used for the classification of Chlorophyta are: type of zoid, mitosis (karyokynesis), cytokinesis, organization level, life cycle, type of gametes, cell wall polysaccharides[20] and more recently genetic data.
Leliaert et al. 2012
Simplified phylogeny of the Chlorophyta, according to Leliaert et al. 2012.[12] Note that many algae previously classified in Chlorophyta are placed here in Streptophyta.
- Viridiplantae
- Chlorophyta
- core chlorophytes
- Ulvophyceae
- Cladophorales
- Dasycladales
- Bryosidales
- Trentepohliales
- Ulvales-Ulotrichales
- Oltmannsiellopsidales
- Chlorophyceae
- Oedogoniales
- Chaetophorales
- Chaetopeltidiales
- Chlamydomonadales
- Sphaeropleales
- Trebouxiophyceae
- Chlorodendrophyceae
- Ulvophyceae
- prasinophytes (paraphyletic)
- Pyramimonadales
- Mamiellophyceae
- Pycnococcaceae
- Nephroselmidophyceae
- Prasinococcales
- Palmophyllales
- core chlorophytes
- Streptophyta
- Chlorophyta
Pombert et al. 2005
A possible classification when Chlorophyta refers to one of the two clades of the Viridiplantae is shown below.[21]
- Class Prasinophyceae T. A. Chr. ex Ø. Moestrup & J. Throndsen
- Class Chlorophyceae Wille
- Class Trebouxiophyceae T. Friedl
- Class Ulvophyceae K. R. Mattox & K. D. Stewart
Lewis & McCourt 2004
- Division Chlorophyta (green algae sensu stricto)
- Subdivision Chlorophytina
- Class Chlorophyceae (chlorophytes)
- Order Chlamydomonadales (+ some Chlorococcales + some Tetrasporales + some Chlorosarcinales)
- Order Sphaeropleales (sensu Deason, plus Bracteacoccus, Schroederia, Scenedesmaceae, Selanastraceae)
- Order Oedogoniales
- Order Chaetopeltidales
- Order Chaetophorales
- Incertae Sedis (Cylindrocapsa clade, Mychonastes clade)
- Class Ulvophyceae (ulvophytes)
- Order Ulotrichales
- Order Ulvales
- Order Siphoncladales/Cladophorales
- Order Caulerpales
- Order Dasycladales
- Class Trebouxiophyceae (trebouxiophytes)
- Order Trebouxiales
- Order Microthamniales
- Order Prasiolales
- Order Chlorellales
- Class Prasinophyceae (prasinophytes)
- Order Pyramimonadales
- Order Mamiellales
- Order Pseudoscourfieldiales
- Order Chlorodendrales
- Incertae sedis (Unnamed clade of coccoid taxa)
- Class Chlorophyceae (chlorophytes)
- Subdivision Chlorophytina
- Division Charophyta (charophyte algae and embryophytes)
- Class Mesostigmatophyceae (mesostigmatophytes)
- Class Chlorokybophyceae (chlorokybophytes)
- Class Klebsormidiophyceae (klebsormidiophytes)
- Class Zygnemophyceae (conjugates)
- Order Zygnematales (filamentous conjugates and saccoderm desmids)
- Order Desmidiales (placoderm desmids)
- Class Coleochaetophyceae (coleochaetophytes)
- Order Coleochaetales
- Subdivision Streptophytina
- Class Charophyceae (reverts to use of GM Smith)
- Order Charales (charophytes sensu stricto)
- Class Embryophyceae (embryophytes)
- Class Charophyceae (reverts to use of GM Smith)
Hoek, Mann and Jahns 1995
Classification of the Chlorophyta, treated as all green algae, according to Hoek, Mann and Jahns 1995.[6]
- Class Prasinophyceae (orders Mamiellales, Pseudocourfeldiales, Pyramimonadales, Chlorodendrales)
- Class Chlorophyceae (orders Volvocales [including the Tetrasporales], Chlorococcales, Chaetophorales, Oedogoniales)
- Class Ulvophyceae (orders Codiolales, Ulvales)
- Class Cladophorophyceae (order Cladophorales)
- Class Bryopsidophyceae (orders Bryopsidales, Halimedales)
- Class Dasycladophyceae (order Dasycladales)
- Class Trentepohliophyceae (order Trentepohliales)
- Class Pleurastrophyceae (order Pleurastrales)
- Incertae sedis (order Prasiolales)
- Class Klebsormidiophyceae (orders Klebsormidiales, Coleochaetales)
- Class Zygnematophyceae (order Zygnematales, Desmidiales)
- Class Charophyceae (order Charales)
In a note added in proof, an alternative classification is presented for the algae of the class Chlorophyceae:
- Class Chlamydophyceae (orders Volvocales, Chlorococcales, Chaetophorales)
- Class Oedogoniophyceae (order Oedogoniales)
- Class Chlorophyceae (order Chlorellales)
Bold and Wynne 1985
Classification of the Chlorophyta and Charophyta according to Bold and Wynne 1985.[22]
- Chlorophyta, Chlorophyceae (16 orders)
- Charophyta, Charophyceae (1 order)
Mattox & Stewart 1984
Classification of the Chlorophyta according to Mattox & Stewart 1984:[23]
- Micromonadophyceae Mattox & Stewart (similar to Prasinophyceae; Tetraselmidiales transferred to Pleurastrophyceae)
- Charophyceae Rabenhorst
- Ulvophyceae Mattox & Stewart
- Pleurastrophyceae Mattox & Stewart
- Tetraselmidiales
- Pleurastrales
- Chlorophyceae Wille in Warming
Fott 1971
Classification of the Chlorophyta according to Fott 1971.[24]
- Class Chlorophyceae
- Order Volvocales
- Order Tetrasporales
- Order Chlorococcales
- Order Ulotrichales
- Suborder Chlorosarcineae
- Suborder Ulotrichineae
- Suborder Oedogoniineae
- Suborder Chaetophorineae
- Order Siphonocladales
- Order Bryopsidales
- Class Conjugatophyceae
- Class Charophyceae
Round 1971
Classification of the Chlorophyta and related algae according to Round 1971.[25]
- "green algae"
- Euglenophyta
- Prasinophyta
- Charophyta
- Chlorophyta
- Zygnemaphyceae (= Conjugatophyceae; orders Mesotaeniales, Zygnematales, Gonatozygales, Desmidiales)
- Oedogoniophyceae (order Oedogoniales)
- Bryopsidophyceae
- Hemisiphoniidae (orders Cladophorales, Sphaeropleales, Acrosiphoniales
- Cystosiphoniidae (orders Dasycladales, Siphonocladales, Chlorochytriales)
- Eusiphoniidae (orders Derbesiales, Codiales, Caulerpales, Dichotomosiphonales, Phyllosiphonales)
- Chlorophyceae
- orders Chlamydomonadales, Volvocales, Polyblepharidales, Tetrasporales, Chlorodendrales, Chlorosarcinales, Chlorococcales
- orders Ulotrichales, Codiolales, Ulvales, Prasiolales, Cylindrocapsales, Microsporales
- orders Chaetophorales, Coleochaetales, Trentepohliales, Pleurococcales, Ulvellales
Smith 1938
Classification of the Chlorophyta according to Smith 1938:
- Class 1. Chlorophyceae
- Order 1. Volvocales
- Family 1. Chlamydomonadaceae
- Family 2. Volvocaceae
- Order 2. Tetrasporales
- Order 3. Ulotrichales
- Family 1. Ulotrichaceae
- Family 2. Microsporaceae
- Family 3. Cylindrocapsaceae
- Family 4. Chaetophoraceae
- Family 5. Protococcaceae
- Family 6. Coleochaetaceae
- Family 7. Trentepohliaceae
- Order 4. Ulvales
- Family 1. Ulvaceae
- Family 2 Schizomeridaceae
- Order 5. Schizogoniales
- Family Schizogoniaceae
- Order 6. Cladophorales
- Family 1. Cladophoraceae
- Family 2. Sphaeropleaceae
- Order 7. Oedogoniales
- Family Oedogoniaceae
- Order 8. Zygnematales
- Family 1. Zygnemataceae
- Family 2. Mesotaeniaceae
- Family 3. Desmidiaceae
- Order 9. Chlorococcales
- Family 1. Chlorococcaceae
- Family 2. Endosphaeraceae
- Family 3. Characiaceae
- Family 4. Protosiphonaceae
- Family 5. Hydrodictyaceae
- Family 6. Oöcystaceae
- Family 7. Scenedesmaceae
- Order 10. Siphonales
- Family 1. Bryopsidaceae
- Family 2. Caulerpaceae
- Family 3. Halicystaceae
- Family 4. Codiaceae
- Family 5. Derbesiaceae
- Family 6. Vaucheriaceae
- Family 7. Phyllosiphonaceae
- Order 11. Siphonocladiales
- Family 1. Valoniaceae
- Family 2. Dasycladaceae
- Class 2. Charophyceae
References
- ↑ Reichenbach, H. G. L. (1828). Conspectus Regni Vegetabilis, p. 23.
- ↑ Pascher A (1914). "Über Flagellaten und Algen". Berichte der deutsche botanischen Gesellschaft 32: 136–160.
- ↑ Adl SM, Simpson AGB, Farmer MA, Andersen RA, Anderson OR, Barta JR, Bowser SS, Brugerolle G, Fensome RA, Fredericq S, James TY, Karpov S, Kugrens P, Krug J, Lane CE, Lewis LA, Lodge J, Lynn DH, Mann DG, McCourt RM, Mendoza L, Moestrup Ø, Mozley-Standridge SE, Nerad TA, Shearer CA, Smirnov AV, Speigel FW, Taylor MFJR (2005). "The new higher level classification of eukaryotes with emphasis on the taxonomy of protists". Journal of Eukaryotic Microbiology 52 (5): 399–451. doi:10.1111/j.1550-7408.2005.00053.x. PMID 16248873.
- 1 2 Guiry, M.D. & Guiry, G.M. (2011). "AlgaeBase : Chlorophyta". World-wide electronic publication, National University of Ireland, Galway. Retrieved 2011-07-26
- ↑ http://archive.org/stream/centuryofprogres00cali#page/126/mode/1up
- 1 2 3 Hoek, C. van den, Mann, D.G. and Jahns, H.M. 1995. Algae An Introduction to Phycology. Cambridge University Press, Cambridge. ISBN 0-521-30419-9
- ↑ "Major Algae Phyla - Table - MSN Encarta". Archived from the original on 2009-10-31.
- ↑ Lewis, Louise A. & McCourt, R.M. (2004). "Green algae and the origin of land plants". Am. J. Bot. 91 (10): 1535–1556. doi:10.3732/ajb.91.10.1535. PMID 21652308
- ↑ Becker, B. & Marin, B. (2009). "Streptophyte algae and the origin of embryophytes". Annals of Botany 103 (7): 999–1004. doi:10.1093/aob/mcp044. PMC 2707909. PMID 19273476
- ↑ Kapraun DF (April 2007). "Nuclear DNA Content Estimates in Green Algal Lineages: Chlorophyta and Streptophyta". Ann. Bot. 99 (4): 677–701. doi:10.1093/aob/mcl294. PMC 2802934. PMID 17272304.
- ↑ Graham LE, Graham JM, Wilcox LW (2009) Algae. 2nd Edition. Benjamin Cummings (Pearson), San Francisco, CA
- 1 2 3 Leliaert, F., Smith, D.R., Moreau, H., Herron, M.D., Verbruggen, H., Delwiche, C.F. & De Clerck, O. (2012). "Phylogeny and molecular evolution of the green algae" (PDF). Critical Reviews in Plant Sciences 31: 1–46. doi:10.1080/07352689.2011.615705.
- ↑ Lewis, Louise; Lewis, Paul (2005). "Unearthing the Molecular Phylodiversity of Desert Soil Green Algae (Chlorophyta)". Systematic Biology 54 (6): 936–947. doi:10.1080/10635150500354852. ISSN 1063-5157.
- ↑ De Wever, A.; Leliaert, F.; Verleyen, E.; Vanormelingen, P.; Van der Gucht, K.; Hodgson, D. A.; Sabbe, K.; Vyverman, W. (2009). "Hidden levels of phylodiversity in Antarctic green algae: further evidence for the existence of glacial refugia". Proceedings of the Royal Society B: Biological Sciences 276 (1673): 3591–3599. doi:10.1098/rspb.2009.0994. ISSN 0962-8452.
- ↑ Leliaert, Frederik; Verbruggen, Heroen; Zechman, Frederick W. (2011). "Into the deep: New discoveries at the base of the green plant phylogeny". BioEssays 33 (9): 683–692. doi:10.1002/bies.201100035. ISSN 0265-9247. PMID 21744372.
- ↑ Lopez-Bautista, J. M. (2006). "Molecular systematics of the subaerial green algal order Trentepohliales: an assessment based on morphological and molecular data". Int J Syst Evol Microbiol 56 (7): 1709–1715. doi:10.1099/ijs.0.63990-0. ISSN 1466-5026.
- ↑ Joubert, J. J. & F. H. J. Rijkenberg (1971). "Parasitic green algae". Annu. Rev. Phytopathol. 9: 45–64. doi:10.1146/annurev.py.09.090171.000401.
- ↑ Nedelcu, Aurora M. (2001). "Complex Patterns of Plastid 16S rRNA Gene Evolution in Nonphotosynthetic Green Algae". Journal of Molecular Evolution 53 (6): 670–679. doi:10.1007/s002390010254. ISSN 0022-2844.
- ↑ Tartar A, Boucias DG, Adams BJ, Becnel JJ (2002). "Phylogenetic analysis identifies the invertebrate pathogen Helicosporidium sp as a green alga (Chlorophyta)". Int J Syst Evol Microbiol 52 (Pt 1): 273–9. PMID 11837312.
- ↑ Lobban, Christopher S.; Wynne, Michael James, eds. (1981). The Biology of Seaweeds. Botanical Monograph Series 17. University of California Press, p. 88.
- ↑ Pombert, Jean-Francxois; et al. (2005). "The Chloroplast Genome Sequence of the Green Alga Pseudendoclonium akinetum (Ulvophyceae) Reveals Unusual Structural Features and New Insights into the Branching Order of Chlorophyte Lineages" (PDF). Mol. Biol. Evol. 22 (9): 1903–1918. doi:10.1093/molbev/msi182. PMID 15930151.
- ↑ Bold, H.C. & Wynne, M.J. (1985). Introduction to the algae : structure and reproduction (2nd ed.). Englewood Cliffs, N.J.: Prentice-Hall. ISBN 978-0-13-477746-7
- ↑ Mattox, K.R. & Stewart, K.D. 1984. Classification of the green algae: a concept based on comparative cytology. Pp. 29‐72, In Irvine, D.E.G. & John, D.M. (eds.), The systematics of Green Algae. The Systematics Association, Special Vol. 27, Academic Press, London.
- ↑ Hoek, C. et al. (1995), p. 483.
- ↑ Round, F.E. (1971). "The taxonomy of the Chlorophyta, 2". Brit. phycol. J. 6 (2): 235–264. doi:10.1080/00071617100650261.
Further reading
- Burrows, E.M. 1991. Seaweeds of the British Isles. Volume 2 Chlorophyta. Natural History Museum, London. ISBN 0-565-00981-8
- Lewis, L. A. & McCourt, R. M. (2004). "Green algae and the origin of land plants". American Journal of Botany 91 (10): 1535–1556. doi:10.3732/ajb.91.10.1535. PMID 21652308.
- Pickett-Heaps, J.D. (1975). Green Algae. Structure, Reproduction and Evolution in Selected Genera. Sinauer Assoc., Stamford, CT; 606 pages.
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